Genetic Basis to Divergence of Sex Pheromones in Two Closely Related Moths, <Emphasis Type="Italic">Ostrinia scapulalis</Emphasis> and <Emphasis Type="Italic">O. zealis</Emphasis>

Crossing experiments between two closely related moths, Ostrinia scapulalis and O. zealis, were conducted to gain insight into the genetic basis of the divergence of female sex pheromones. The sex pheromone of O. scapulalis comprises (E)-11- and (Z)-11-tetradecenyl acetates (E11 and Z11), and distinct genetic variation is found in the blend of components. This variation is largely controlled by a single autosomal locus with two alleles, A^E(sca) and A^Z(sca). E-type (A^E(sca)A^E(sca)) females produce a pheromone with amean E11:Z11 ratio of 99:1, whereas Z-type (A^Z(sca)A^Z(sca)) and I-type (A^E(sca)A^Z(sca)) females produce a pheromone with a mean of 3:97 and 64:36, respectively. O. zealis is distinctive in that it has a third pheromone component, (Z)-9-tetradecenyl acetate (Z9), in addition to E11 and Z11, and the typical blend ratio is 60:35:5 (Z9:E11:Z11). Our study revealed that Z9 production in O. zealis is mainly regulated by an autosomal recessive gene phr^(zea), which is suggested to be involved in the chain-shortening of a pheromone precursor fatty acid, and linked to A^E(zea), a gene corresponding to A^E(sca) in O. scapulalis. A few mutations in a gene involved in pheromone production could explain the dramatic shift between a two-component pheromone communication system in O. scapulalis and a three-component system in O. zealis.     

Potential for evolution of resistance to pheromones: Interindividual and interpopulational variation in chemical communication system of pink bollworm moth

After an extensive examination of the release rates and blend ratios of pheromonal components emitted by field-collected femalePectinophora gossypiella (Saunders), we find no evidence of resistance to pheromones applied to cotton fields to disrupt mating. Females from fields with 3–5 years of exposure to disruptant pheromones as well as those from fields with only minimal exposure to disruptant pheromones emitted (Z,Z)-7,11-hexadecadienyl acetate at a rate of ca. 0.1 ng/min and (Z,E)7,11-hexadecadienyl acetate at ca. 0.06 ng/min. The ratio of pheromonal components was much less variable than the measured emission rate and was centered about a 61:39Z, Z to Z,E ratio. In contrast to the blend ratio emitted by females, the composition of the pheromonal blend used in monitoring populations and disrupting mating is centered about 50:50 Z,Z to Z.E. In general there was a remarkable consistency in the release rate and blend ratio among populations of females throughout southern California and those from a laboratory colony. It would appear that, although resistance to theP. gossypiella pheromone is still a very real possibility when it is used heavily in pest management as a mating disruptant, there are current agricultural practices and conditions which would hinder its development. Pectinophora gossypiella (Saunders), Lepidoptera: Gelechiidae.     

Potential for Use of Synthetic Sex Pheromone for Mating Disruption of the Olive Pyralid Moth, Euzophera pinguis

The potential for pheromone-based mating disruption of the olive pyralid moth (OPM), Euzophera pinguis, in olive groves was investigated during the second flight period in small-plot trials in 2002. The female of this species emits a blend of (9Z,12E)-tetradecadien-1-ol and (9Z,12E)-tetradecadienyl acetate, which were synthesized for field tests. Mating disruption efficacy in 0.8-ha trials was evaluated using two parameters: reduction of male capture in pheromone traps and reduction of infestation in infestation-prone sites. White rubber septa containing 10 mg of pheromone blend as disruptant were applied at a density of 50 septa/ha for each treatment. Mean catches of E. pinguis males in pheromone traps were greatly reduced (> 95%) in pheromone-treated plots relative to similar traps placed in control plots. In addition, significant reductions were recorded (35-40%) in the oviposition and infestation levels during pheromone treatment. The total amount of pheromone blend released from disruption dispensers during the field trials was estimated to average 5.4 mg/ha/day, over 56 days.     

Female Sex Pheromone Polymorphism in Adzuki Bean Borer, Ostrinia scapulalis, Is Similar to That in European Corn Borer, O. nubilalis

Individual analysis of the female sex pheromone of the adzuki bean borer, Ostrinia scapulalis, has shown that the sex pheromone of this species comprised (E)-11-tetradecenyl acetate (E11–14:OAc) and (Z)-11-tetradecenyl acetate (Z11–14:OAc) at variable blend ratios. The pheromone blend could be tentatively categorized into three types with respect to the proportion of E11–14:OAc: E type (94–100%, median 99.2%), Z type (0–16%, median 3.0%), and intermediate type (I type, 48–85%, median 63.7%). In addition to the identity of components, the blend ratios in the three types were similar to those of the E strain, Z strain, and hybrid of the European corn borer, O. nubilalis, respectively. This finding suggests that two closely related but morphologically distinct species, O. scapulalis and O. nubilalis, share almost the same sex pheromone communication systems. The significance of this similarity in the two sibling species is discussed.     

Heritable Variation in the Sex Pheromone of the Almond Moth, Cadra cautella

The sex pheromone of the female almond moth, Cadra cautella, constitutes a highly variable signal. Age and period of the L/D cycle had significant effects on the variation in titers of (Z,E)-9,12-tetradecadienyl acetate (Z9,E12-14:Ac) and (Z)-9-tetradecenyl acetate (Z9-14:Ac) and the ratio of Z9,E12-14:Ac to Z9-14:Ac. Across age classes, minimum and maximum mean titers (ng +/- SE) of Z9,E12-14:Ac and Z9-14:Ac ranged from 8.7 +/- 0.46 to 21 +/- 1.0 and from 2.0 +/- 0.14 to 2.9 +/- 0.15, respectively; the mean ratio of Z9,E12-14:Ac to Z9-14:Ac ranged from 5.2 +/- 0.21 to 11 +/- 0.40. The titers of Z9,E12-14:Ac and Z9-14:Ac and the ratio of Z9,E12-14:Ac to Z9-14:Ac were highest at the onset of the scotophase and lowest at the onset of the photophase. Similarly, the titer of Z9,E12-14:Ac and the ratio of Z9,E12-14:Ac to Z9-14:Ac were highest in females sampled 1 d postemergence and declined over all age classes. The titer of Z9-14:Ac increased from day 1 to day 2, and then declined to levels equivalent to day 1. Analysis of pupal and adult mass demonstrated a positive correlation (r = 0.874, P < 0.001); however, the titer of neither Z9,E12-14:Ac or Z9-14:Ac nor the ratio of Z9,E12-14:Ac to Z9-14:Ac were significantly correlated with pupal mass. Age-related variations in pheromone titer and ratios were heritable. The narrow-sense heritability of the observed variation in 1-d-old females, 1 hr into the scotophase is 1.2 +/- 0.32, 0.75 +/- 0.24, and 0.46 +/- 0.17 for the titer of Z9,E12-14:Ac and Z9-14:Ac and the ratio of Z9,E12-14:Ac to Z9-14:Ac, respectively. A significant additive genetic correlation was observed between that of Z9,E12-14:Ac and Z9-14:Ac, but not between that of either Z9,E12-14:Ac or Z9-14:Ac and the ratio of Z9,E12-14:Ac to Z9-14:Ac. The results of the genetic analyses suggest that the traits of titer of Z9,E12-14:Ac and Z9-14:Ac have a greater potential to respond to selection than the trait of ratio of Z9,E12-14:Ac to Z9-14:Ac.     

Major Sex Pheromone Components of the Australian Gum Leaf Skeletonizer Uraba lugens: (10E,12Z)-Hexadecadien-1-yl Acetate and (10E,12Z)-Hexadecadien-1-ol

Two sex pheromone components of the gum leaf skeletonizer, Uraba lugens (Lepidoptera: Nolidae), recently established in New Zealand, were identified. Gas chromatography (GC) electroantennographic detection analyses of female pheromone gland extracts gave three compounds that consistently elicited antennal responses. Chemical analyses, using GC and GC-mass spectrometry, in conjunction with 4-methyl-1,2,4-triazoline-3,5-dione and dimethyldisulfide derivatizations, identified these compounds as (10E,12Z)-hexadecadien-1-yl acetate (E10,Z12-16:Ac), (10E,12Z)-hexadecadien-1-ol (E10,Z12-16:OH), and (Z)-11-hexadecen-1-yl acetate (Z11-16:Ac). A trapping trial in Queensland, Australia, in 2002, indicated that a blend of the two major components E10,Z12-16:Ac and E10,Z12-16:OH could attract gum leaf skeletonizer males. In the same trial, E10,Z12-16:Ac alone trapped large numbers of an unidentified nolid, Nola spp. Further trials in Auckland, New Zealand established that these two components were sufficient and necessary for trap catch of males; adding minor gland components, (10E,12E)-hexadecadien-1-yl acetate (E10,E12-16:Ac), Z11-16:Ac, or octadecan-1-ol (18:OH), to the two-component lure did not result in increased trap catches. Behavioral observations and gland analyses of the Auckland population revealed that female moths begin calling soon after emergence, with peak calling and pheromone production occurring 7 hr into the scotophase. Analysis of gland extract at two-hourly intervals during the first activity period showed that the ratio of E10,Z12-16:Ac to E10,Z12-16:OH (mean of 86: 14, respectively) and pheromone titer were fairly constant. No qualitative or quantitative differences in pheromone components were detected between gland extracts from Tasmanian univoltine and Auckland bivoltine populations of U. lugens.     

Sex Pheromones and Reproductive Behavior of Spodoptera litura (Fabricius) Moths Reared from Larvae Treated with Four Insecticides

When Spodoptera litura (Fabricius) larvae were treated with the insecticides deltamethrin, endosulfan, malathion, and carbaryl at dosages causing 30% mortality (LD30), female moths reared from survivors showed similar patterns in the frequency of calling behavior. However, females reared from larvae dosed with deltamethrin had significantly higher titers of sex pheromone than those treated with endosulfan, malathion, or carbaryl, or control individuals. The ratio of the sex pheromone blend, Z9,E12-14:OAc to Z9,E11-14:OAc in females from the deltamethrin treatment group differed significantly from that of female moths in other treatments. In a wind-tunnel bioassay, males reared from larvae treated with deltamethrin, malathion, or carbaryl were less likely to display behaviors resulting in the location of a sex pheromone source than males reared from larvae treated with endosulfan or a solvent control (acetone). Mating success was significantly decreased in pairs containing either males or females reared from larvae that survived treatment with deltamethrin.     

Heritable variation in pheromone response of the pink bollworm,Pectinophora gossypiella (Lepidoptera: Gelechiidae)

Heritability of variation in male pheromone response by pink bollworm moths,Pectinophora gossypiella (Saunders), was examined using a still-air, wing-fanning bioassay. Heritability (±SE) of overall responsiveness, as measured by the mean duration of wing fanning to the blend of pheromone components produced by females [4456 ratio of (Z, E)- to (Z, Z)-7,11-hexadecadienyl acetate], was 0.385 ± 0.095. Heritabilities of wingfanning duration to blends with 25 and 65%Z, E isomer were 0.377 ± 0.113 and –0.145 ± 0.103, respectively. These findings indicate an asymmetry in the genetic component of variation in response to pheromone blends with high and low proportions of theZ, E isomer. An index of response specificity for individual males was developed based on the response to an off-blend (either 25 or 65%Z, E isomer) relative to the response to the 44%Z, E blend. Heritabilities of response specificity were 0.117 ± 0.059 and –0.043 ± 0.067 for the 25 and 65%Z, E blends, respectively.     

Potential for evolution of resistance to pheromones

FemalePectinophora gossypiella (Saunders) from most of the desert cotton-growing areas of southern California emitted significantly more pheromone in 1984 and 1985 than in preceding years (1982 and 1983). This increase amounted to almost 20% by 1985. It is unlikely that this small change would represent effective resistance to disruptant pheromones, but this increase could reflect the result of selection pressure imposed by the use of mating disruption for population control. A worldwide survey of emitted pheromone from this species found that there was much more variation in the emission rate than the blend ratio of the two pheromone components. The emitted blend ratio was remarkably consistent over time (in southern California) and throughout the worldwide range of the insect. Small differences in the blend ratio that were detected probably have no major biological significance because of the relatively broad response spectrum of males to changes in the blend of pheromonal components. Populations of males did not consist of several phenotypes, each with a different preference for specific blend ratios. Rather, the broad response spectrum to blend ratios in a population can be attributed to variation in the response of any individual. Therefore, selection for a response phenotype that is narrowly tuned to the blend emitted by females may be difficult.     

Sex Pheromone of the Dogwood Borer, <Emphasis Type="Italic">Synanthedon scitula</Emphasis>

The sex pheromone of female dogwood borers (DWB) Synanthedon scitula (Harris) (Lepidoptera: Sesiidae) was determined to be an 88:6:6 ternary blend of (Z,Z)-3,13-octadecadienyl acetate (Z,Z-3,13-ODDA), (E,Z)-2,13-octadecadienyl acetate (E,Z-2,13-ODDA), and (Z,E)-3,13-octadecadienyl acetate (Z,E-3,13-ODDA) by gas chromatography–electroantennographic detection (GC–EAD) and gas chromatography–mass spectrometry (GC–MS). The major sex pheromone component, Z,Z-3,13-ODDA, was attractive as a single component. A blend of Z,Z-3,13-ODDA with 1–3% of E,Z-2,13-ODDA (binary blend) was more attractive than the single component. A third component, Z,E-3,13-ODDA, was sometimes observed in GC–EAD analyses, and enhanced attraction to the binary blend in some field bioassays. Lures containing 1 mg of binary and ternary blends attracted 18 and 28 times more male DWB moths, respectively, than caged virgin females in field trials. Attraction was strongly antagonized by addition of as little as 0.5% of E,Z-3,13-octadecadienyl acetate (E,Z-3,13-ODDA). In a period of 12 wk in 2004, more than 60,000 males were captured in sticky traps baited with synthetic pheromone blends in six apple orchards in Virginia, West Virginia, and North Carolina. Lure longevity trials showed that ∼76% of the pheromone remained in rubber septum lures after 12 wk in the field.     

Genetic aspects of interpopulational differences in pheromone blend of cabbage looper moth,Trichoplusia ni

The genetic basis of interpopulational differences in the pheromone blend emitted by the cabbage looper moth,Trichoplusia ni (Hübner), was examined by crossing individuals from a field-derived population (P₁) with individuals from a long-maintained laboratory colony (P₂). These colonies differed in the emission rate and relative proportions of four of the five known minor pheromone components, but not in the emission rate of the major component, (Z)-7-dodecenyl acetate (Z7-12Ac). These differences in pheromone blend were quantitatively small but biologically significant, because in the field, males responded preferentially to traps baited with a pheromone blend that is similar to that emitted by P₁ females relative to a blend similar to that emitted by P₂ females. In initial crosses, variation in the quantity and quality of pheromone blends among families of P₁, P₂, and F₁ hybrid females was examined. In F₁ females the relative proportions (quantity relative to the major component) of (Z)-5-dodecenyl acetate (Z5-12Ac) and (Z)-7-tetradecenyl acetate (Z7-14Ac) were intermediate to parental lines. In a second more extensive set of crosses, analyses included P₁, P₂, F₁, F₂, and selected backcrosses. The relative proportion of Z5-12Ac, Z7-14Ac, and Z9-14Ac emitted by F₁ females were intermediate to parental lines. The frequency distributions of relative proportions of these components emitted by females were not consistent with those expected under a single autosomal or sex-linked gene hypothesis, suggesting that more than one gene is involved in the quantitative differences in the pheromone blend.     

Evaluation of the Major Female Eurytoma amygdali Sex Pheromone Components, (Z,Z)-6,9-Tricosadiene and (Z,Z)-6,9-Pentacosadiene for Male Attraction in Field Tests

We evaluated the attraction of male almond seed wasp Eurytoma amygdali to the synthetic alkadienes (Z,Z)-6,9-tricosadiene and (Z,Z)-6,9-pentacosadiene and their blend in almond orchards using baited rubber septa attached to cardboard rectangular adhesive traps. The two alkadienes were recently isolated from virgin female whole body extracts and SPME collected volatiles. The alkenes (Z)-9-tricosene, (Z)-9-pentacosene, and (Z)-9-heptacosene, present in female extracts, were also added to the blend of the alkadienes and tested. The alkadienes tested individually attracted males when the traps were baited with doses ranging from 10 to 30 mg/trap. The maximum number of males was attracted to traps baited with 10 mg of a (Z,Z)-6,9-C(23:2):(Z,Z)-6,9-C(25:2) blend at a ratio of 7:3. Results with the three alkenes added to the blend were inconclusive because of low populations. The present study on E. amygdali is the first one reporting attraction of males to synthetic sex pheromone components in field trials for a Eurytomidae species. The synthetic alkadienes blend offers the potential to develop an effective system for monitoring populations of the almond seed wasp in almond orchards.     

(Z)-11-octadecenyl acetate,an aggregation pheromone inDrosophila simulans

Existence of a male-produced pheromone, which attracts both males and females in a wind-tunnel olfactometer, has been demonstrated inDrosophila simulans (Sturtevant). A pheromone component was identified as (Z)-11-octadecenyl acetate (Z11-18: Ac), also known ascis-vaccenyl acetate. The pheromone is synergized by food volatiles. In bioassay ca. 1/1000 of a mature male equivalent of Z11-18: Ac is attractive and activity increases with increased amounts of Z11-18: Ac. Flies do not begin responding to Z11-18: Ac until after they have been away from food for at least 2 hr. Z11-18: Ac is transferred from the male to the female during mating, and the female emits the majority of the transferred Z11-18: Ac within 6 hr after mating.     

Variability in pheromone composition and periodicity of pheromone titer in potato tuberworm moth,Phthorimaea operculella (Lepidoptera: Gelechiidae)

The ratios and quantities of the pheromone components, (E,Z)-4,7-tridecadien-1-yl acetate (diene) and (E,Z,Z)-4,7.10-tridecatrien-1-yl acetate (triene), in the glands of individual female potato tuberworm moths (Phthorimaea operculella) originating from the United States (California) and Japan (Nagoya) were analyzed by gas chromatography. Quantities of glandextracted pheromone components of Nagoya females fluctuated in a periodic fashion during the photoperiod. Maximal titers coincided with the onset of scotophase (and calling), then gradually declined to minimal levels soon after lights-on. The average daily pheromone quantities decreased significantly as females aged. Both populations exhibited considerable variation in the ratio of the two components. The proportions of triene in the blend ranged from 27% to 88% (triene –X = 56 ± 13% SD; CV = 23%) for California females and from 16% to 71% (42 ± 13%; CV = 31%) for Nagoya females. Nagoya females also stored significantly higher amounts of pheromone in their glands (8.6 ± 3.9 ng) than did California females (2.7 ± 1.4). The differences between the populations, while substantial, would probably not be sufficient to impart a barrier to panmixis, given the wide range of component ratios favored by the males.     

(<Emphasis Type="Italic">Z,Z</Emphasis>)-6,9-Heneicosadien-11-One: Major Sex Pheromone Component Of Painted Apple Moth, <Emphasis Type="Italic">Teia anartoides</Emphasis>

(Z, Z)-6,9-Heneicosadien-11-one (Z6Z9-11-one-21Hy) was identified as the major sex pheromone component of the painted apple moth (PAM), Teia anartoides (Lepidoptera: Lymantriidae), on the basis of (1) comparative gas chromatographic-electroantennographic detection (GC-EAD) analyses, GC-mass spectrometry (MS), high-performance liquid chromatography (HPLC)-MS, and HPLC-UV/visible spectroscopy of pheromone gland extracts and authentic standards; (2) GC-EAD analyses of effluvia of calling females; and (3) wind tunnel and field trapping experiments with a synthetic standard. In field experiments in Australia, synthetic Z6Z9-11-one-21Hy as a single component attracted male moths. Wind tunnel experiments suggested that a 4-component blend consisting of Z6Z9-11-one-21Hy, (6Z,9R,10S)-cis-9,10-epoxy-heneicosene (Z6-9R10S-epo-21Hy), (E, E)-7,9-heneicosadien-6,11-dione (E7E9-6,11-dione-21Hy), and 6-hydroxy-(E, E)-7,9-heneicosadien-11-one (E7E9-6-ol-11-one-21Hy) (all present in pheromone gland extracts) might induce more males to orient toward, approach, and contact the source than did Z6Z9-11-one-21Hy as a single component. Additional experiments are needed to determine conclusively whether or not Z6-9R10S-epo-21Hy, E7E9-6,11-dione-21Hy, and E7E9-6-ol-11-one-21Hy might be minor sex pheromone components of PAM. Moreover, attractiveness of synthetic pheromone and virgin PAM females needs to be compared to determine whether synthetic pheromone could replace PAM females as trap baits in the program to monitor eradication of exotic PAM in New Zealand.     

EFFECT OF PBAN ON PHEROMONE PRODUCTION BY MATED <Emphasis Type="Italic">Heliothis virescens</Emphasis> AND <Emphasis Type="Italic">Heliothis subflexa</Emphasis> FEMALES

Mated female Heliothis virescens and H. subflexa were induced to produce sex pheromone during the photophase by injection of pheromone biosynthesis activating neuropeptide (PBAN). When injected with 1 pmol Hez-PBAN, the total amount of pheromone that could be extracted from glands of mated females during the photophase was similar to that extracted from virgin females in the scotophase. The PBAN-induced profile of pheromone components was compared between mated, PBAN-injected females and virgin females during spring and fall. Virgin females exhibited some differences in the relative composition of the pheromone blend between spring and fall, but no such temporal differences were detected in PBAN-injected, mated females. Because the temporal variation in pheromone blend composition was greater for virgin females than for PBAN-injected females, PBAN can be used to determine a females native pheromone phenotype. This procedure has the advantages that pheromone glands can be extracted during the photophase, from mated females that have already oviposited.     

Mating Disruption of Citrus Leafminer Mediated by a Noncompetitive Mechanism at a Remarkably Low Pheromone Release Rate

The citrus leafminer, Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae), is a worldwide pest of citrus. A season-long investigation was conducted that evaluated mating disruption for this pest. Effective disruption of the male P. citrella orientation to pheromone traps (98%) and reduced flush infestation by larvae was achieved for 221 d with two deployments of a 3:1 blend of (Z,Z,E)-7,11,13-hexadecatrienal/(Z,Z)-7,11-hexadecadienal at a remarkably low rate of 1.5 g active ingredient (AI)/ha per deployment. To gain insight into the mechanism that mediates the disruption of P. citrella, male moth catch was quantified in replicated plots of citrus treated with varying densities of pheromone dispensers. The densities of septum dispensers compared were: 0 (0/ha, 0.0 g AI/ha), 0.2 (one every fifth tree or 35/ha, 0.05 g AI/ha), 1 (215/ha, 0.29 g AI/ha), and 5 per tree (1,100/ha, 1.5 g AI/ha). Profile analysis by previously published mathematical methods matched predictions of noncompetitive mating disruption. Behavioral observations of male P. citrella in the field revealed that males did not approach mating disruption dispensers in any of the dispenser density treatments. The current report presents the first set of profile analyses combined with direct behavioral observations consistent with previously published theoretical predictions for a noncompetitive mechanism of mating disruption. The results suggest that disruption of P. citrella should be effective even at high population densities given the density-independent nature of disruption for this species and the remarkably low rate of pheromone per hectare required for efficacy.     

Heritable variation of sex pheromone composition and the potential for evolution of resistance to pheromone-based control of the Indian meal moth,Plodia interpunctella

The short-term evolutionary effect of pheromone-based mating disruption on the mating ability of the Indian meal moth, Plodia interpunctella, was investigated. Three independent selection lines were established, and the mating ability of moths in plastic tents treated with high doses of pheromone and in control tents was compared for two consecutive generations. In addition, the heritability of the sex pheromone blend, measured as the ratio of two major pheromone components (Z,E)-9,12-tetradecadienyl acetate and (Z,E)-9,12-tetradecadienol, was estimated. Based on a mother–daughter regression analysis including 21 families, the heritability of the pheromone blend was 0.65 ± 0.14, indicating a potential for evolutionary change of the character. However, no increase in mating ability of females in pheromone-treated tents or alteration of the pheromone blend was observed in any selection line when compared with control lines, indicating no or weak selection on the pheromone blend as well as other traits influencing mating ability of this species under the created mating disruption conditions. Factors contributing to the lack of selection effects are discussed.     

Identification and synthesis of a male-produced sex pheromone from the stink bug Chlorochroa sayi

The reproductive behavior of the stink bug Chlorochroa sayiwas studied in the laboratory. There was a sexual maturation period of about 10 days before bugs began mating. Sexually mature adult bugs engaged in courtship consisting of antennation and head-butting of the female by the male, before the female adopted a receptive posture and copulation occurred. Both sexes mated multiple times during their life-spans, with the mean duration of copulations of virgin bugs (42.3 ± 19.6 min) and experienced bugs (37.3 ± 28.4 min) being similar. Most matings were initiated in the late afternoon or evening, when pheromone production by males was greatest. Males transferred sperm and nutrients constituting about 17% of their body weight to females during mating. Three male-specific components, methyl geranate, methyl citronellate, and methyl (E)-6-2,3-dihydrofarnesoate in a ratio of 100 : 0.45 : 1.6, were first detected in volatiles collected from male bugs on green beans about 9–12 days after the final molt to the adult stage. In vertical Y-tube bioassays, females were attracted to odors from mature male bugs, and to a blend of the three male-produced components. Low numbers of females also were attracted in field trials with the three-component blend. The relatively weak attraction may be a result of other, as yet unknown cues being required in addition to the pheromone, such as visual or substrate-borne vibrational cues.     

Individual variation in sex pheromone of smaller tea tortrix moth,Adoxophyes sp. (Lepidoptera: Tortricidae)

Female smaller tea tortrix mothsAdoxophyes sp. (Lepidoptera: Tortricidae), which initiated calling at 1, 2, or 3 days old, respectively, were analyzed individually for (Z)-11-tetradecenyl acetate (Z11-14:OAc) and (Z)-9-tetradecenyl acetate (Z9–14: OAc) in the pheromone gland via GLC. Among different age groups, broad and similar distributions were found for pheromone quantity (¯X=58.6±52.9 ng/female; range 1.3–219.8 ng/female). The ratio of the two pheromone components averaged 6535 but ranged from 8416 to 4060. The significance of the pheromone blend variation to the attraction of males was tested in a field experiment. The ratio of males trapped by the most attractive blend versus the least attractive one was 2.2.